We measured gas exchange and various leaf parameters of ash (Fraxinus angustifolia Vahl.) and oak (Quercus robur L.) in the high canopy and of lime (Tilia cordata Mill.) in the lower canopy of a planted, 120-year-old floodplain forest in southern Moravia, Czech Republic. The high-canopy leaves of F. angustifolia and Q. robur had nitrogen concentrations on a leaf area basis (N(area)) that were twice those of low-canopy leaves of T. cordata. Upper-canopy leaves of F. angustifolia had a photosynthetic rate at light saturation (A(max)) of about 16 micromol CO2 m(-2) s(-1), whereas A(max) of the upper-canopy foliage of Q. robur achieved only about two thirds of this value. Contrary to previous investigations of photosynthetic performance in monospecific stands, leaves of the uppermost branches of T. cordata at 15-m height had the highest A(max) and transpiration rate among the species studied. Water-use efficiency (WUE) was low in T. cordata at 15-m canopy height, whereas WUE was significantly higher for Q. robur leaves at 27-m height than for the other species. Leaves of T. cordata at 15-m height showed the strongest relationship between A(max) and N(area) (R2 = 0.90) followed by F. angustifolia (R2 = 0.69). The strong correlation between photosynthesis and nitrogen concentration in T. cordata at 15 m, together with the steep regression slope for the A(max):N(area) relationship, indicated that nitrogen allocation to the photosynthetic apparatus resulted in high nitrogen-use efficiency of light-saturated photosynthesis (PNUE). Despite differences in PNUE among species, PNUE was fairly constant for leaves sampled from the same canopy position, suggesting that single-leaf parameters are matched to optimize PNUE for prevailing light conditions. High PNUE in T. cordata at 15 m partially compensated for the species' subordinate position in the canopy, and may be an important mechanism for its coexistence in highly structured vegetation.

Department of Systematic Botany and Ecology Universität Ulm Germany

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